Turkish Journal of Fisheries and Aquatic Sciences 8: 125-132 (2008)

Life- History Traits and Decadal Trends in the Growth Parameters of Golden putitora (Hamilton 1822) from the Himalayan Stretch of the Ganga River System Prakah Nautiyal1,*, Anees Fatma Rizvi2, P. Dhasmanaa1 1 H N B Garhwal University, Department of Zoology, Srinagar, 246174, . 2 Institute of Applied Sciences, Allahabad 211002, India.

* Corresponding Author: Tel.: -; Fax: -; Received 07 August 2006 E-mail: [email protected]; ribbonfish123@ rediffmail.com Accepted 12 March 2008 Abstract

Life-history traits such as age and body growth patterns of (Hamilton 1822), inhabiting the lesser Himalayan stretch of the Ganga river system in the Garhwal region were examined during 1994 and 1995. The maximum age was recorded to 17+ whereas harvestable size was 65.6 cm. The asymptotic length (L∞) exhibited a relative decline from 272 -1 -1 cm (K = 0.035 year ) in 1980-81 to 216 cm (K = 0.041 year ) in 1994-95 resulting in the corresponding increase of the growth coefficient (K). The total, natural and fishing mortality coefficients also exhibited a similar pattern 0.366, 0.054 and 0.312 and 0.58, 0.063, 0.517 year-1, respectively during 1980-81 and 1994-95. This was reflected in the exploitation rate and ratio also, 0.376 and 0.852 and 0.7 and 0.891 year-1 respectively for 1980-81 and 1994-95

Key words: age, growth, asymptotic length, mortality, exploitation, decadal trends.

Introduction descend as the floods subside. T. putitora thus exhibits a tri-phased migration (Nautiyal and Lal, Desai (2003) has listed 10 valid of 1984; Nautiyal, 2002). mahseer from , India, China and Southeast Among the various species of Mahseer in India Asia, though more species have been listed recently T. putitora, familiarly known as the Golden or (Annexure 1). Dwivedi (2002) is of the opinion that Himalayan mahseer attains the largest size, 275 cm (9 the members of this group are found in rivers in the ft) in length and 54 kg (118 lb) in weight (Talwar and northern Plains of India and extend towards west Jhingran, 1991; Wikipedia, 2006). The size was through the river Indus in Pakistan and Euphrates and recorded by Hamilton in 1822. The first author (PN) Tigris in Iraq and to north in China. A number of recorded a size of 137.7 cm in 1980-81. Owing to its species are found in the southeast also (Kiat, 2004). size, golden colour, beautiful appearance and flavour, Annexure 2 lists the countries and their locations the fish is exploited thereby constituting an important compatible with distributional range of the Golden fishery along the Himalayan foothills. Exploitation as mahseer. It has been introduced in the southeast as far well as other factors (Nautiyal, 1984, 1989, 1990, as New Papua Guinea. Mahseer is a spectacular game 1994) has led to a decline in numbers in the fish and it constitutes main fishery in the Sivalik Himalayan stretch of the Ganga. The following Himalaya and uplands of the Deccan Plateau. Three observation supports this view, “Qasim and Qayyum species of the genus Tor occur in the Himalaya, Tor (1961) studied the breeding biology of T. putitora putitora, T. tor and T. progenius, the former is from the Ganga near Aligarh (far downstream of the prevalent in Jammu and Kashmir, Himachal Pradesh, foothills at Hardwar). Today, it is restricted to the Uttarakhand, and even the northeast, while foothills (Rishikesh-Hardwar).” In the face of high latter two are restricted to Central and East Himalaya. intraspecific competition, especially due to high The Himalayan species live in foothill section density (density-dependent) it seems reasonable for (Sivaliks) of glacierfed Ganga where waters are not individuals to disperse in the vicinity of ideal habitat. ice-cold. The species of Peninsular India inhabit the Large numbers would facilitate dispersal while rivers devoid of glaciers. Mahseer is known migrate decline would restrict the population to the most ideal into small tributaries during the breeding season part of the habitat. Two factors seem to be causing the (Beavan, 1877; Desai, 2003). For the Himalayan decline, over fishing and habitat degradation – mahseer first flooding caused by snowmelt is a signal fragmentation. Hence, when (prior to 1960’s) the for upstream migration into the glacierfed tributaries. Himalayan mahseer occurred in large numbers they The migrants remain in them till the onset of monsoon were able to populate the then Ganga till Narora near when the brooders again ascend into the flooded Aligarh (the last limits of the upper stretch of the spring fed streams for breeding while others Ganga) far downstream of the foothills. But as they (juveniles and adolescents) descend to the foothills. became few they got restricted to the foothills only. The brooders and new recruits (year-old juveniles) Decline has been observed in most of the Indo-

© Central Fisheries Research Institute (CFRI) Trabzon, Turkey and Japan International Cooperation Agency (JICA)

126 P. Nautiyal et al. / Turk. J. Fish. Aquat. Sci. 8: 125-132 (2008)

Gangetic drainages over the west (Sehgal et al., 1971; exploitation rate and ratio of this species. The Joshi, 1988; Sunder and Joshi 1977; Sehgal, 1994) information is considered a prerequisite for managing and central Himalaya (Shrestha, 1997). the fishery and conservation in case of threatened fish The mahseer are considered as ‘endangered’ species. Hence, investigations were undertaken on (Khan and Sinha, 2000). The decline of the mahseer age, growth parameters, mortality, exploitation rate (T. putitora and T. tor) population in the Lake and ratio. Nainital was mentioned by Raj (1945). Das and Aloka (1978) found biological indicators of pollution in Study Area Lake Nainital and related it to the depletion of Mahseer fish (T. putitora). Das and Upadhyay (1979) All specimens of T. putitora were obtained studied the qualitative and quantitative fluctuations of downstream of Ajeetpur (altitude 273 m; 29°52'50'' N; plankton in two Kumaon lakes, Nainital and Bhimtal 78°10'23'' E) from the Ganga, Raiwala (altitude 340 to determine the causes of the failure of the Mahseer m; 30°3'18'' N; 78°13'54" E) from the Saung, Banghat (T. tor and T. putitora) fisheries in the Kumaon lakes. (altitude 560 m; 29°57'12" N; 78°13'23" E) from the The ‘endangered’ status of the Himalayan mahseer Nayar and Srinagar (altitude 550 m; 30°13'30"; population, however, is debatable as there are still 78°49'39" E) from the Alaknanda (Figure 1). some pockets where this species predominates. The adult population inhabits the foothill stretch Mahseer has kept a steady profile in terms of average of the Ganga and migrates upstream into the Saung size in the reservoirs. For the last 22 years the average and Nayar for spawning. Therefore, these two weight has ranged between 1.2 to 1.6 kg while the tributaries harbour the younger stages and were total landings have fluctuated between 10 to 102 sampled to obtain them. Beside breeding (Sehgal, tonnes during 1995-96 to 1997-98 and the average 1972) there may be other reasons for which the fish weight of mahseer was 1.2 kg (Himachal Pradesh may migrate upstream which may involve factors like Fisheries Department, 2007). The Himalayan mahseer maintaining food supply, homing instinct and learning has been largely studied for natural history, status and by reinforcing memory (Nautiyal et al., 2001). This conservation practices in India and Nepal (Nautiyal, can be explained by the presence of migrants of 1994; Shrestha, 1997). Barring recent attempts to varying sizes (10-137 cm, juveniles, adolescents both investigate the population biology (Bhatt et al., immature, maturing virgins and mature adults) in the 1998a; 1998b; 2000) much remains to be known Alaknanda for 3-4 months (February to June), about the dynamics of T. putitora. FAO has produced whereas the Nayar and Saung largely invite the estimates of population parameters from Nepal brooders (Figure 2). In fact the large size (>80 cm) (Fishbase, 2003). There is no account of mortality and can be obtained only during the migratory phase and

Figure 1. The study area of the Gangetic drainage in Himalaya.

P. Nautiyal et al. / Turk. J. Fish. Aquat. Sci. 8: 125-132 (2008) 127 were not found in the foothill stretch harbouring the read the ‘key scales’ of T. putitora, obtained from the major part of the mahseer population in the mountain dorsal fin region above lateral line. The annulus section of the Ganga (Nautiyal, 2002). formation was determined according to the criterion suggested by Begenal and Tesch (1978). Materials and Methods Data Analysis Field Sampling The length frequency data obtained at monthly

Random samples were obtained from the fish intervals during 1980-81 and 1994-95 were pooled markets, landings sold off locally by fishermen or and finally raised to the annual total catch of the from those procured by the contractor at these species. The growth parameters L∞ and K were stations. At all sites of collection, T. putitora were estimated using the Gulland and Holt (1959) formula: landed using gillnets (7-10 cm mesh size), cast nets -∆L / ∆t = K*L , and seines. Hook and line was the most effective ∞ method of catching adults. For each fish, total length was recorded to the nearest cm and weight in grams. whereas to was calculated by the von The fish ranged from 4.9 to 137.7 cm during 1980-81 Bertalanffy’s plot. (sample size (n) = 132) and from 4.0 to 135.9 cm The growth parameters computed were used to during 1994-95 (n = 815). Length at age frequency determine the natural mortality (M) using the methods of Srinath (1998) 1.532*K, where K is the growth data obtained during 1980-81 and 1994-95 were grouped into size classes with 10 cm interval length in coefficient. The total mortality (Z) was determined by order to have adequate number of fish in each size using the Beverton and Holt (1956) method, class for growth study (Table 1). L - L Age and growth were computed from the data Z = K ∞ c generated by using Carl Zeiss Jena Documenter to Lc - Lc

Ganga in foothills 1 FEEDING GROUNDS OF THE JUVENILES , 2 3 ADOLESCENTS AND THE ADULTS [October-March] 1, 2 & 3 1 & 2 1&2 lower himalayan stretch of the Ganga [March-June]

3 mid-himalayan rivers 1 Alaknanda, Bhagirathi [March-June] 1 & 2 3 3 3 3

1 & 2

Saung like spring -fed streams near foothills [1&2, March -June; 3, July - September]

Nayar like springfed streams in mid -himalaya [1&2; March-June; 3, July-September]

SPAWNING GROUNDS OF ADULTS, REARING & FEEDING GROUNDS OF LARVAL AND POST LARVAL STAGES

Figure 2. Adult population of Himalayan Mahaseer Tor putitora migrates form foothill stretch of the Ganga to upstream of the Saung and Nayar for breeding and maintaining food supply etc.

128 P. Nautiyal et al. / Turk. J. Fish. Aquat. Sci. 8: 125-132 (2008)

where Lc is the length at which 50% of the fish 1994-95 differed primarily with respect to L∞ while entering the gear are retained and Lc is the average the growth coefficient was 0.035 for 1981-82 and length of the entire catch. 0.041 for 1994-95 (Figure 3 and 4) during respective

The Fishing mortality (F) was estimated by years. The to was calculated as 0.031 for year 1980-81 subtracting M by Z value obtained according to the and 0.0153 year for 1994-95. The von Bertalanffy’s Beverton and Holt (1956) method. Exploitation rate (1938) equation for growth in length for this species (U) and ratio (E) were obtained by the Beverton and could thus be written as, – 0.055 (t- 0.031) Holt (1957) formula: Lt = 272.2 (1- e ) (1980-81) Lt = 216 (1- e – 0.056 (t- 0.015)) (1994-95) U= F/Z (1-e -Z) and E= F/Z. The instantaneous rate of the total mortality Results coefficient (Z) was estimated to be 0.366 per year in 1980-81 while 0.58 per year in 1994-95. Similarly, Age of 0+ to 17+ years was determined from the the natural mortality coefficient (M) estimates were scales of Tor putitora population. The largest 0.054 per year for 1980-81 and 0.063 per year for specimen of this species was measured as 137.7 and 1994-95. The fishing mortality (F) estimated was 135.9 cm for 1980-81 and 1994-95, respectively, the 0.312 for 1980-81 and 0.517 for 1994-95. The estimated age from length at age frequency data being estimated exploitation ratio (E) and rate (U) were 17.5 years (Table 1). Growth parameters obtained by 0.852 and 0.376 for 1980-81, and 0.891 and 0.7 for the Gulland and Holt plot (1959) for 1980-81 and 1994-95.

Table 1. Length frequency key for studying growth parameters of T. putitora during 1980-81 and 1994-95.

Size interval 1980-81 1994-95 (cm) mean length (cm) mean age frequency mean length (cm) mean age frequency 1-10 6.54 0.5 26 6.60 0.5 147 11-20 14 1.5 28 14.17 1.5 381 21-30 23.83 2.5 15 25.87 2.5 116 31-40 35.9 3.5 13 36.65 3.6 56 41-50 44.11 4.5 10 46.27 4.59 43 51-60 56.85 5.5 8 56.85 5.5 23 61-70 68.01 6.5 7 67 6.5 28 71-80 78.03 7.41 5 77.08 8.5 13 81-90 88.92 8.43 4 85.5 9.59 2 91-100 97.53 9.25 4 95.6 10.51 1 101-110 107.02 10.26 4 106.3 11.51 1 111-120 115.44 12.5 4 115.3 13.7 2 121-130 129 16.5 3 126.5 16.5 1 131-140 137.7 17.5 1 135.9 17.5 1

14 11 K=-b 12 10 9 10

t 8

Δ t

8 Δ 7 L / Δ L / 6 6 Δ 5 K=-b 4 4 2 3 0 5 10 15 2 L 051015 L

Figure 3. Gulland and Holt plot (1980-81) showing the length Figure 4. Gulland and Holt plot (1994-95) showing the increment or growth rate ΔL/Δt in y-axis is plotted against length increment or growth rate ΔL/Δt in Y-axis is plotted mean length during the corresponding year. The regression against mean length during the corresponding year X-axis. line given intercept (a) = 9.637 and slope (b) = -0.0354 and the The regression line given intercept (a) = 8.8699 and slope intersection point between the regression line in x-axis given (b)=-0.0409 and the intersection point between the L ∞. regression line in x-axis given L∞.

P. Nautiyal et al. / Turk. J. Fish. Aquat. Sci. 8: 125-132 (2008) 129

Discussion years, it seems to be under alarmingly high fishing pressure, which has damaged its fishery (Nautiyal et Stock identification is important because al., 1998; Bhatt et al., 2004). continuous fishing on a particular population, if The commercially exploited fish species seem to homogenous, has a direct effect on the population of exhibit decline in the asymptotic length (L∞) and may the same species in every other locality. Conversely, hence be related to excessive exploitation. T. tor the where a species consists of two or more stocks, first cousin of T. putitora, as both sometime occur in fishing at any one locality, whatever its magnitude has the same rivers, performed better growth especially in no effect on the other unfished stock of the same the Yamuna basin rivers (northeast India). The species. The stock of T. putitora from the mountain estimates for T. tor; L∞ = 787, 822 and 946 mm; K = stretch of the Gangetic system has been demonstrated 0.61, 0.78, 0.50 year-1; F = 2.9, 3.44, 4.57 year-1 in the to be homogeneous (Nautiyal and Lal, 1988; Bhatt et Paisuni, Ken and Tons, respectively, indicate low L∞ al., 1998a; 1998b), implying the need to determine the in rivers with high fishing mortality. Similarly, the features of the stock. The growth parameters have total mortality (Z) was quite high for T. tor (4.08 year- been determined from the actual data for the first time 1 to 5.57 year-1) if compared with L. rohita (3.2 year-1 for this species. to 4.19 year-1) and L. calbasu (0.98 year-1 to 1.86 year- 1) in these rivers indicating greater fishing pressure on Population Dynamics T. tor and L. rohita, as both are highly important and priced fish like T. putitora, L. calbasu being least Johal and Kingra (1989) using the Walford priced (Dwivedi, Unpublished). method obtained an estimate of 135 cm L∞ for T. The natural mortality coefficient (M) of a fish is putitora from the Gobindsagar, which was quite directly related to the growth coefficient ‘K’ and smaller then for the Gangetic stock. The harvestable inversely related to the asymptotic length (L∞) and the size was computed to be 65 cm (Nautiyal, 2006). The lifespan (Beverton and Holt, 1956). The natural maximum attainable size computed for Cirrhinus mortality of T. putitora was relatively low (M = 0.054 mrigala (90 cm), Catla catla (120 cm), Labeo rohita -1 -1 year , K = 0.035 year in 1980-81 compared to M = (85 cm), Cyprinus carpio (95 cm) and 0.063 year-1, K=0.041 year-1 in 1994-95). The Hypophthalmichthys molitrix (110 cm) are quite low mortality rate has thus increased. Among the three (Johal and Tandon, 1987a; 1987b; Bhandari et al., components of mortality (total, fishing, natural) 1993) suggesting that the Himalayan mahseer has a computed in the present study, mortality was mainly longer life span. The Amur carp (common carp) is due to fishing which points to heavy fishing pressure known to attain a size of 90 cm and an age of 16 years on them. This was also reflected in the exploitation (Nikolskii, 1980). rates and ratios, 0.376 and 0.852 and 0.7 and 0.891 Beverton and Holt (1957) pointed out that the year-1 respectively for 1980-81 and 1994-95. two parameters of growth; asymptotic length (L∞) and growth coefficient (K) are inversely proportional to Conclusions each other. It implies that fishes with high L∞ should have lower K values and vis-a-vis. T. putitora, a cold T. putitora, inhabiting the mountain rivers, is a water inhabitant, with maximum observed sizes (Lmax) game and food fish with long life span. The of 137.7 cm and 135.9 cm, respectively in 1980-81 Himalayan mahseer exhibited slow growth -1 and 1994-95, had L∞ = 272 cm and K = 0.035 year comparatively to its first cousin T. tor that is habiting -1 compared with L∞ of 216 cm and K of 0.041 year for in tropical waters. Analysis of the decadal trends respective years. Since T. putitora sample for 1980-81 revealed that asymptotic length (L∞) decreased from had a slightly larger size than 1994-95 it had 1980-81 (272 cm), to 1994-95 (216 cm). An relatively higher L∞ and slightly lower K. Thus, the increasing pattern was obtained for total, natural and -1 - present estimate of asymptotic length and growth fishing mortality coefficients 0.366 year , 0.054 year coefficient for T. putitora are justified. Growth 1 and 0.312 year-1 and 0.58 year-1, 0.063 year-1, 0.517 -1 parameters computed earlier (L∞ = 275 cm, K = 0.070 year , respectively during 1980-81 and 1994-95. -1 year , to = 0.25 year) for T. putitora were slightly Increased mortality rate during the 1994-95 higher than the present estimate. Since then such a mentioned above be a sign of over exploitation rate size has not been reported for the Himalayan mahseer. and ratio 0.7 year-1 and 0.891 year-1 respectively -1 -1 A female measuring 148.0 cm from the Sarju River, compared to 0.376 year and 0.852 year respectively Kumaun Himalaya India is the only report available for 1980-81.Thus there is a need to manage the over last two decades. A size less than that was fishery of T. putitora to prevent a collapse of both the reported in the early eighties (Nautiyal and Lal, fishery and the population. 1981). The present sample with a still lesser length -1 Acknowledgement had lower K = 0.041 year and to= 0.015 year. The L∞

= 272 cm in 1980-81 and 216 cm in 1994-95 indicated tendency of decrease in size, the age groups The authors are thankful to Dr. P. Gaur being 17+ during respective years. With increasing Professor and Head and Professor H. R. Singh former exploitation ratio and fishing mortality in subsequent Head, Department of Zoology, University of

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Allahabad for providing all the research facilities. We Dwivedi A.C. 2006. Age Structure of Commercially also thank Dr. S. K. Chakraborty (Principal Scientist) Exploited Fish Stocks of Ganga River System (Banda CIFE (Mumbai) for providing invaluable suggestions. -Mirzapur section). PhD. thesis Allahabad: University of Allahabad, Dwivedi, S.N. 2002. “Mahseer” the of India- References conservation in . In: K.K. Vass, H.S. Raina (Eds.), Highland Fish. Aqua. Res. Man. Bagenal, T.B. and Tesch, F.W. 1978. Age and growth. In: National Research Centre on Coldwater Fisheries T.B. Bagenal (Ed.), Method for Assessment fish (ICAR) Bhimtal: 30-37. rd Production in Fresh Waters 3 Edition Blackwell Fishbase, 2003. www.fishbase.org/manual/Keyfacts.htm Scientific Publication., Oxford: 101- 136. (accessed 15.07.2003) Beavan, R. 1877. Handbook of freshwater fishes of India. Gulland, J.A. and Holt, S.J. 1959. 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Annexure I. Scientific names where genus equals Tor (Adopted from FishBase) Scientific Name Author Valid Name English Name Tor arabicus (Trewawas, 1941) Barbus arabicus Tor ater Roberts, 1999 Tor ater Tor blanci (Pellegrin & Fang, 1940) Neolissochilus blanci Tor brevifilis (Peters, 1881) Tor brevifilis Tor brevifilis brevifilis (Peters, 1881) Tor brevifilis Tor brevifilis hainanensis Wu, 1977 Tor brevifilis Tor canis (Valenciennes, 1842) Barbus canis Tor chelynoides (McClelland, 1839) Naziritor chelynoides Dark mahseer Tor douronensis (Valenciennes, 1842) Tor douronensis River carp Tor hamiltoni Gray, 1834 Tor tor Mahseer Tor hemispinus Chen & Chu, 1985 Tor hemispinus Tor khudree (Sykes, 1839) Tor khudree Deccan mahseer Tor khudree longispinnis (Günther, 1868) Tor khudree Deccan mahseer Tor khudree malabaricus Jerdon, 1849 Tor khudree Deccan mahseer Tor kulkarni Menon, 1992 Tor kulkarnii Tor kulkarnii Menon, 1992 Tor kulkarnii Tor laterivittatus Zhou & Cui, 1996 Tor laterivittatus Tor longipinnis (Weber & de Beaufort, 1916) Neolissochilus longipinnis Tor manningsi (de Beaufort , 1933) Barbus nanningsi Tor mosal (Hamilton, 1822) Tor putitora Putitor mahseer Tor mosal mahanadicus David, 1953 Tor khudree Deccan mahseer Tor mussullah (Sykes, 1839) Tor mussullah High-backed mahseer Tor musullah (Sykes, 1839) Tor mussullah High-backed mahseer Tor nedgia (Rüppell, 1836) Barbus nedgia

132 P. Nautiyal et al. / Turk. J. Fish. Aquat. Sci. 8: 125-132 (2008)

Annexure I. (Continued)

Scientific Name Author Valid Name English Name Tor polylepis Zhou & Cui, 1996 Tor polylepis Tor progeneius (McClelland, 1839) Tor progeneius Jungha mahseer Tor putitora (Hamilton, 1822) Tor putitora Putitor mahseer Tor qiaojiensis Wu, 1977 Tor qiaojiensis

Tor reinii (Günther, 1874) Barbus reinii Tor sinensis Wu, 1977 Tor sinensis Tor soro (Valenciennes, 1842) Tor

Annexure I. Occurrence records of Tor putitora (+ Compatible with distributional range; * requires matching against distributional range). (source FishBase, The original information has been modified by grouping catalog no,. for locations visited by one author)

Country Year Collector Identifier, Catalog No. Information _ 1940 _ BMNH 1940.3.25.1 Foothillo of Himalayas _ 1964 Meehean, O. L. USNM 00257749 Meehean, O. L., W. PAKISTAN, 16 MI. E. OF PINDI China 1995 Kullander, S O & S.O. Kullander/ 1997 Kullander, S O & F Fang, P R CHINA, Yunnan, Main F Fang NRM 33258; 36115, 36116 Mengla County, Ying Jiang County+ India 1889 _ BMNH 1889.10.29.18-19 Dehra Dun, North West Provinces+ India 1889 _ BMNH 1889.10.29.16-17 Dehra Dun, North West Provinces + India 1880 McClelland BMNH 1880.2.2.5 McClelland, River of India + India 1870 BMNH 1870.5.18.15 Malabar +

India 1868 Day, F. BMNH 1868.10.27.18 Day, F., Bowany+ India 1843 BMNH 1843.2.25.29 Gumnah, Sehamapore+ India 1986 BMNH 1986.11.6.1 Almorah,+ India 1944 BMNH 1944.7.31.4 Ihelum, River Ihelum, India + India 1932 BMNH 1932.2.20.11 Nagrota, Punjab + India no year A. W. Herre. CAS 134609 A. W. Herre., Nandhaur River. + India no year A. W. HERRE CAS 133965 A. W. HERRE, Dehra Dun, Eastern Doons; Uttar Pradesh, India + India 1934 _ BMNH 1934.10.17.37 Bengal + India 1932 _ BMNH 1932.2.20.13 Ravi, Ravi River, Madhopur, Punjab + 1932.2.20.12 India 1954 Menon, A.G.K. BMNH 1954.5.20.7-8 Sarda, Menon, A.G.K., Sarda River, Tanakpur, Naital, United Provinces + Myanmar no year Malaise, R S.O. Kullander/1997 Malaise, R, MYANMAR, Bago Division, Kachin NRM 31609 31871, 31607 State, Bago Division * Myanmar 1893 _ BMNH 1893.6.30.31-40 Nampandet Myanmar 1935 Maung Lu Daw S.O. Kullander/1997 Maung Lu Daw, MYANMAR, Sagaing Division NRM 18794 * Myanmar 1997 Fang, F & A S.O. Kullander/ 1997 Fang, F & A Roos, MYANMAR, Kachin State * Roos NRM 36309 Myanmar 1934 Malaise, R S.O. Kullander/1997 NRM Malaise, R, MYANMAR, Shan State * 10407 Nepal no year A.C. Taft H. DeWitt/1959, CAS 152925 A.C. Taft, Near Pokhra (Pokhara); purchased +

H. DeWitt/1959 CAS 152924 A.C. Taft, Inlet stream to Phewa Tal Lake, near Nepal no year A.C. Taft Pokhra + Nepal 1996 David Edds David Edds David Edds, At Mulghat, on road from Dharan to KU 29597 29619 29535, Hile, Kachali river confluence, Kahare, Sabha 29530, 29520, 28668, 29033, river confluence, 1 hour walk south of Tu, Just 29537, 29458, 28806 29007, east of Tumlingtar, Brahamadev, Andhi Mohan - 29470 29119 29436, 29079, Andhi river confluence, Piluwaa river confluence, 29059, 29348, 29412 29484, Gorangi - about 4 km west of Chisapani, Just 28999 downstream from irrigation project along Raj, Khalte, Chapang, Purchased at Koshi barrage, Just east of Katasi, Khairenitar, Narayangarh, first feeder creek bridge south of Chatra on ro, Purchased at Chisapani , Kharkhareghat , Nimaa + Pakistan 1908 BMNH 1908.12.28.101 Lahore Sri Lanka 1929 BMNH 1929.7.2.5 Beira Lake, Colombo, Ceylon*